Habitat Selection by Black Kite Breeders and Floaters
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Habitat selection by Black kite breeders and floaters: Implications for conservation management of raptor floaters ⇑ Alessandro Tanferna, Lidia López-Jiménez, Julio Blas, Fernando Hiraldo, Fabrizio Sergio Department of Conservation Biology, Estación Biológica de Doñana, Consejo Superior de Investigación Científicas, Sevilla 41092, Spain abstract Preserving large predators is important but challenging because these species are typically wide-ranging, select multiple habitats at different scales and often present spatial or habitat separation between the breeder and floater sectors of a population. In addition, most of our knowledge on raptor floaters’ habitat requirements comes from large solitary species, whose floaters often occupy temporary settlement areas spatially separate from breeding locations. Here, we examine space and habitat use by a loosely colonial, wetland-dependent raptor, the Black kiteMilvus ( migrans), in a population where floaters co-exist with Keywords: territory holders, enabling a direct comparison of their habitat preferences. The study was conducted Habitat selection in Doñana National Park (South-Western Spain), a seasonally drying marshland currently surrounded Black kite Milvus migrans by intensive agriculture and rice-fields. Intensive radio-tracking revealed that breeders and floaters Radio-tracking selected and avoided the same habitats despite a radical, four-to-eight fold difference in their home- Habitat restoration range dimensions: all kites over-selected open habitats suitable for their aerial foraging modes and Non-breeders avoided woodland and farmland. These results suggest a continuum of raptor population structures rang- ing from solitary species whose floaters select different habitats than breeders and are concentrated in spatially separate settlement areas, to colonial and semi-social species whose floaters fully coexist with breeders with shared habitat preferences. Both extremes of this continuum will pose challenges for con- servation management. In solitary species, special conservation efforts may be required to identify and manage temporary settlement areas, while in gregarious species, the larger ranges of floaters may expose them to different threats than breeders, whose occurrence and consequences may be subtle to identify. 1. Introduction (e.g. David Smith, 1993; Ferrer and Harte, 1997; Crabtree and Shel- don, 1999; Balbontín, 2005). The latter adds complexity to strate- Preserving and managing large vertebrate predators is becom-gic management targeting long-term population persistence, ing increasingly important as a way to maintain high levels especialof bio-ly because non-breeding animals are difficult to study diversity (Estes et al., 2011), but poses special challengesdue for to their cryptic behaviour, differential habitat selection, spatial several reasons. First, these species are characterised by largeseparation from breeders, or potential long-distance dispersal (e.g. home-ranges, which cannot be easily encompassed within pro-Zack and Stutchbury, 1992; Rohner, 1997; Whitfield et al., 2009a; tected areas (e.g. Newton, 1979; Clark et al., 1999; Ray Penterianiet al., et al., 2011). As a result, there is little knowledge on the 2005). Second, they frequently select habitat features at multipledifferences in habitat choices between the breeders and floaters of scales, from the micro-scale to the landscape-level (Sánchez-Zapaa tapopulation, and conservation planning is often biased to protect and Calvo, 1999; Thompson and McGarigal, 2002; Ciarniello etthe al., habitats preferred by the breeding sector of predator popula- 2007), which requires broad-level management plans (e.g. tions (e.g. Real and Mañosa, 1996; Whitfield et al., 2006a). Also, Whitfield et al., 2006a). Third, they may use different habitatsmost atof the (scarce) available knowledge is heavily biased towards different times of the year (Boal et al., 2005; Schmitz largeet al.,species of solitary birds of prey, whose floaters are typically 2010). Finally, the populations of large predatory vertebrates concentare rated in so-called ‘‘temporary settlement areas’’, where frequently composed of a sector of territorial breeders, often theycon- select different habitats than breeders (Ferrer and Harte, centrated in resource-rich sites, and a sector of non-breeding 1997;indi- Balbontín, 2005; Caro et al., 2011; Penteriani et al., 2011). viduals, frequently located far away from the breeding groundsAs a result, little is known of smaller species with different social systems, such as colonial or loosely colonial species. ⇑ Corresponding author. Tel.: +34 605 482244. Because of all the above, there is a high need for further multi- E-mail addresses:[email protected], [email protected] (A. Tanfer- scale habitat selection studies on both breeding and non-breeding na). individuals of predatory vertebrates, particularly of gregarious inantor conspecifics (Sergio et al., 2009; Blas and Hiraldo, 2010; Blas semi-gregarious species. Here we provide such a study by examin-et al., 2011). Sexual role division during reproduction follows the ing the space and habitat requirements of the breeders and floatersusual scheme for raptors (Newton, 1979): the male provides most of a semi-social raptor, the Black Milvuskite (migrans). In particu- of the prey for the female and offspring while the female performs lar, we use data from a 3-year radio-tracking study on themost Black of the incubation, brooding and nest guarding. kite population of Doñana National Park (South-Western Spain), one of the most renowned and biodiversity-rich reserves of Europe.2.2. Study area Our study system is a good model for the goals presented above due to several reasons. (1) The Black kite is a semi-gregarious Therap- study was conducted in Doñana National Park, located tor, which in Doñana mainly breeds in loose colonies (Sergiowithin et al., the estuary of the Guadalquivir river, along the coast of 2005). (2) In this population, floaters coexist with breeders the(BlasAtlantic Ocean in South-Western Spain (6°120–6°400W, et al., 2009; Sergio et al., 2009, 2011a). This allows the study36°48 0–37of °200N). The five main macro-habitats observed in the habitat selection by the two status categories while controllingpark include: (1) seasonally drying marshland (hereafter ‘‘marsh- for differences in habitat availability, avoiding the problem of land’’),com- (2) Mediterranean scrubland or grassland with scattered paring the decisions by groups of individuals occupying separatecork oaks Quercus( suber) (hereafter ‘‘dehesa’’), (3) extensive scrub- areas characterised by different landscapes. (3) Doñana Nationalland on sandy soil (hereafter ’’scrubland’’), a mixture of different Park is an island of semi-natural vegetation subject to dynamicdegradation stages of autochthonous Mediterranean scrubland habitat management and transformation. Outside the park, drain-(Castroviejo, 1993), including patches dominatedPistacia by lentis- age of the seasonal marshes in the second half of the 20thcus centuryand Myrtus communisor byHalimium halimifolium, Ulex spp., has generated a matrix of intensive farmland, dominated byStauraca rice nthus genistoidesand Erica spp.; (4) mobile sand dunes fields to the north-east, whose suitability for wildlife species alongis lar- the ocean coast, and (5) extensive forests of stonePinus pine gely unknown. Inside the protected area, all habitats are tradition-pinea and smaller woodlots dominated by Cork oaksEucalyptus or ally actively managed (e.g. Fernández-Delgado, 2005). For spp. trees (Castroviejo, 1993). A mosaic of intensively cultivated example, large patches of forest have been recently thinned landsor re- and rice fields surrounds the park. moved, while the hydrology of the seasonal marshes that charac- terise the park is subjected to a recently implemented large-scale2.3. Field methods restoration program (Project ‘‘Doñana 2005’’: García Novo and Marín Cabrera, 2005a), which included the restoration of variousBetween 2007 and 2009 we trapped 38 Black kites by cannon- sites totalling more than 502 ofkm seasonal marshland which netting (Fig. 1) and equipped them with a conventional backpack had been originally converted to agriculture (García Novo transmitterand (TW-3 of 15 g; life expectancy = 1.4 years; Biotrack Marín Cabrera, 2005b; Santamaría et al., 2005; Martín-LópezLtd., Wareham Dorset, UK), which was fitted with a Teflon harness et al., 2011). The above described habitat changes and active( Kenward,man- 2001). The sex, status and sampling period of tracked agement inside and outside the park call for more solid knowledgekites are specified in Table 1. Kites were monitored every 3–4 days of the habitat preferences of key species such as Black kites,and which all locations, obtained by triangulation, were GIS mapped are the most abundant large predators in the park and whichthrough de- the software ArcView 3.2 (ArcView GIS, Redlands, CA, pend heavily on woodland for nesting and marshland for huntingUSA). In each tracking day, all marked kites were searched simul- (Sergio et al., 2011b). Understanding habitat preferences of taneouslykey while driving along a network of paved and dirt roads indicator species could be fundamental to forecast future impactscovering the entire park and its surroundings, thus sampling areas of habitat management and to implement more efficient post-both close and far from nest concentrations. Also, the starting point intervention monitoring. and sequence of survey roads were varied each time, in order to